Accumulation of Phenylpropanoids in the Cotyledons of Morning Glory (Pharbitis nil) Seedlings during the Induction of Flowering by Low Temperature Treatment, and the Effect of Precedent Exposure to High-Intensity Light

Extracts from the cotyledons of seedlings of Pharbitis nil strain‘Violet’ cultured at low temperature, which inducestheir flowering even in continuous light, with or without precedentexposure to high-intensity light, which shortens the periodof low temperature required for flowering, were analyzed byHPLC for substances correlating with the flower-inducing process.The content of two phenylpropanoids were found to increase duringthe low-temperature, and were identified as 3-O-feruloylquinicacid and dehydrodiconiferyl alcohol-13-O-ß-D-glucoside.The increase was more rapid in the cotyledons exposed to high-intensitylight before the low-temperature. This suggests that the accumulationof these compounds is correlated to the promotive effect ofhigh-intensity light on the flower-induction by low temperature. (Received March 7, 1994; Accepted April 2, 1994)     

Increase in the Activity of Phenylalanine Ammonia-Lyase during the Non-Photoperiodic Induction of Flowering in Seedlings of Morning Glory (Pharbitis nil)

The activity of phenylalanine ammonia-lyase in the seedlingsof Pharbitis nil strains ‘Violet’ and ‘Kidachi’during the non-photoperiodic induction of flowering increasedin close association with the accumulation of phenylpropanoidsand the induction of flower buds. This observation suggestedthat the increased activity of phenyalanine ammonia-lyase causedthe accumulation of the phenylpropanoids and that the activationof the biosynthesis of phenylpropanoids might be involved inthe non-photoperiodic induction of flowering in the seedlingsof ‘Violet’ and ‘Kidachi’. (Received September 6, 1994; Accepted December 15, 1994)     

Effect of High-intensity Light Given Prior to Low-temperature Treatment on the Long-day Flowering of Pharbitis nil

Pharbitis nil, strain Violet which had been exposed to high-intensitylight (18,000 lux at 23?C) for 7 days followed by a low-temperaturetreatment (13–14?C) for 7 days initiated flower buds evenunder continuous light, but plants given these treatments inreverse order failed to bud. Three days of high-intensity lightat 23?C was most effective in promoting the flower-inducingeffect of the subsequent low-temperature period. Six days oflow temperature following the 3-day high-intensity light periodinduced near-maximum flowering response. DCMU (5?10^–6M) given during the high-intensity light period inhibited flowering,but when given during or after the low-temperature period itwas ineffective. DCMU at the same concentration given before,during or after an inductive 16-hr dark period at 26?C did notinhibit flowering. Sucrose, ATP, NADPH and some other reducingagents tested did not nullify the DCMU effect nor substitutefor the effect of high-intensity light. But, the high-intensitylight effect could be substituted, at least partly, by 5-chlorosalicylicacid, 3,4-dichlorobenzoic acid and some other benzoic acid derivatives,which are highly effective in inducing long-day flowering inthe short-day plant, Lemna paucicostata. (Received October 20, 1981; Accepted February 3, 1982)     

Primary and Secondary Induction Requirements for Flowering of Contrasting European Varieties of Lolium perenne

The flowering requirements of six European varieties of Loliumperenne L. were studied in controlled environments. In experimentson primary induction, flowering was recorded after transferto long days (LD) in a greenhouse at 12–24°C. In experimentson secondary induction, primary induction was first accomplishedat 6°C/10 h daylength for 12 weeks. When evaluated by the50% heading criterion, the requirement for duration of primaryinduction at 6°C/8 h daylength was <3 weeks in Mediterranean,5–6 weeks in Central European and 7–8 weeks in Scandinavianvarieties. While ‘Veyo’ (Italy) flowered profuselyregardless of temperature or daylength during primary induction,critical temperatures for primary induction in SD and LD were15 and 11°C in ‘Baca’ (Czech Republic) and 11and 7°C in ‘Falster’ (Denmark). The criticalphotoperiod for secondary induction at 15°C ranged from12 h in ‘Veyo’ and 14 h in ‘Baca’ to16.5 h in ‘Falster’ and 17.5 in ‘Kleppe’(Norway). The critical number of LD cycles varied correspondingly.While the Central and North European varieties required fewerLD cycles for 50% heading at 18 than at 12°C, ‘Veyo’showed the opposite response. It is concluded that the requirementsfor both primary and secondary induction of Lolium perenne increasewith increasing latitude of origin of the germplasm. In oneexperiment, 39–87% of the inflorescences came from tillersthat were not visible on transfer from primary to secondaryinduction, thus it is also concluded that there is no juvenilestage in tillers of Lolium perenne. Copyright 2000 Annals ofBotany Company Daylength, flowering, juvenility, perennial ryegrass (Lolium perenne L.), primary induction, secondary induction, temperature, varieties, vernalization     

Participation of polyamines in the flowering of the short-day plant Pharbitis nil

The effect of the exogenous application of polyamines on the flowering induction of the short-day plant Pharbtis nil was investigated. Putrescine, spermidine and spermine applied on the cotyledons of 4-day seedlings had no significant effect on the flowering of this plant under conditions of full induction caused by a 16-hour-long inductive night. Under the conditions of partial induction caused by a 13-hour-long subinductive night, polyamines inhibit or stimulate flowering, depending on the time of application. Also, inhibitors of the biosynthesis of polyamines influenced the flowering process. Analysis of endogenous polyamines revealed significant fluctuations in their content in cotyledons during an inductive night, as well as under continuous light conditions. Particularly large changes occurred in spermidine and spermine levels. The putrescine level in induced seedlings was lower than in non-induced ones. However, induced seedlings contained a higher level of spermine and spermidine. The highest spermidine and spermine levels were observed at the 8th h of the night, although the total concentration of spermine during photoinduction was always 2–3 times lower than that of spermidine. A break in the inductive night, leading to a complete inhibition of flowering, had caused significant changes in the polyamine level by the end of the night. The results suggest that the flowering induction of Pharbitis nil took place at a low putrescine level and increased spermidine and spermine levels.     

Effects of Monochromatic Radiations on Growth of Pelargonium Callus Tissue

Pith callus tissues were grown under continuous blue (450 mµ),green (545 mµ), red (650 mµ), and ‘white’(full-spectrum) light, and in the dark for 22 days at 27±2°C at energy levels of 15,000 ergs cm^–2 sec^–1. Mean increases in fresh weight of tissues grown under ‘white’and blue light were significantly greater than those of tissuesgrown in green and red light and in the dark. Tissues grownin the dark yielded mean fresh weight increases significantlylower than tissues grown under blue, red, and ‘white’light. No significant differences were shown between blue and‘white’, red and green, and green and dark treatmentsrespectively. Cell differentiation occurred in all treatmentsonly to the extent of vessel element formation. There were nodifferences in degree of differentiation between treatments. It was proposed that the high-energy reaction of photomorphogenesiswas in operation in the Pelargonium callus tissue. The resultsindicated the presence in the tissue of high-energy photoreceptor(s).The use of high-intensity, incandescent illumination for experimentalprocedures approximating natural conditions of irradiation wasindicated as desirable for pith callus tissues of Pelargoniumzonale var. Enchantress Fiat.     

Evidence for the Involvement of Calcium Ions in the Photoperiodic Induction of Flowering in Pharbitis nil

The effects of ethyleneglycol-bis-(ß-aminoethyl ether)N,N,N',N'-tetraacetic acid, a specific chelator of Ca²⁺ ions;lanthanum chloride, a calcium channel blocker; and chlorpromazine,a calmodulin antagonist, on the photoperiodic induction of floweringin Pharbitis nil were studied by perfusing the plants with aqueoussolutions of the various compounds. All these compounds inhibitedflowering when applied before an inductive 16-h dark periodbut they did not inhibit flowering when applied after the inductivedark period. The results imply that Ca²⁺ ions are involved inthe photoperiodic induction of flowering in P. nil. (Received August 14, 1992; Accepted November 24, 1992)     

Trade Winds Reduce Growth and Influence Gas Exchange Patterns in Papaya Seedlings

Four experiments were conducted to determine the effect of tradewinds in Guam, USA, on growth and gas exchange of three papaya(Carica papaya L.) cultivars. ‘Known You 1’, ‘Sunrise’,and ‘Tainung 2’ papaya seedlings at two differentstages of development were exposed to 0, 36 or 100% ambientwind. Wind exposure reduced stem height and leaf or stem dryweight in most cases, but had little influence on root growth.Net CO₂assimilation (A_CO2) at midday was lower for seedlingsexposed to wind than for those protected from wind. Dark respirationof exposed seedlings increased as much as 120% above that ofthe protected seedlings during the night. Wind exposure decreasedwhole plant evapotranspiration by up to 36% during the photoperiod,but increased evapotranspiration by 58–87% during thenocturnal period. Responses to wind exposure were similar amongcultivars, except that growth of ‘Tainung 2’ seedlingswas less affected by wind than that of the other cultivars.Seedlings that were exposed to the various wind treatments fromgermination onwards were less influenced by wind exposure thanwere seedlings that were grown in a protected nursery beforebeing exposed to the various wind treatments. These data indicatethat: (1) ambient trade winds in Guam are strong enough to decreasethe growth of papaya seedlings; (2) plant age influences theresponse; (3) stem and leaf growth are more influenced thanroot growth; and (4) decreasedA_CO2 and increased dark respirationmay be partly responsible for growth reduction. Copyright 2001Annals of Botany Company Carica papaya, gas exchange, wind     

The Importance of Light Intensity for Pollen Tube Growth and Embryo Survival in Wheat x Maize Crosses

The success of Triticum aestivumxZea mays crosses, used to producewheat doubled haploids, is influenced by light intensity. Toexamine the basis for this response, pollen tube growth, embryosurvival and indicators of photosynthetic rate were measuredin two wheat cultivars (‘Karamu’ and ‘Kotuku’)crossed with maize at two irradiance levels (250 or 750 µmolm^-2s^-1, PAR). Pollen tube growth was significantly affectedby light intensity in ‘Karamu’ plants but not in‘Kotuku’ plants, despite both cultivars being pollinatedby the same maize source. The percentage of pollen tubes reachingthe cavity between the ovarian wall and integuments, or in themicropyle of ‘Karamu’ plants at high light intensity(65%) was nearly three-times greater than that at low lightintensity (22%). Thus, either low light intensity can affectthe maternal wheat plant in a way that inhibits pollen tubegrowth and/or high light intensity may promote pollen tube growthin ‘Karamu’ plants. Significant differences in ratesof electron transport in plants grown at the two light intensitiesindicated that the rate of photosynthesis may also have an effecton pollen tube growth. These results have importance for improvingthe efficiency of wheat x maize crosses and other wide cerealcrosses. Copyright 2001 Annals of Botany Company Intergeneric hybridization, light intensity, pollen tube growth, embryo survival, Triticum aestivum, wheat,Zea mays , maize     

Accumulation of Cytokinin-Induced Betacyanin in Specific Cells of Amaranthus tricolor Seedlings

The accumulation of betacyanin, in dark-grown Amaranthus tricolorseedlings, in response to cytokinins or red light, occurs mainlyin two specific tissues, the lower epidermal cells of the cotyledons(with the exception of guard cells), and the endodermis of thehypocotyl. The possible significance of this ‘spatialpattern of competence’ is discussed, together with theconcept of target cells in relation to plant hormones. The effect of removing exogenously supplied cytokinin at varioustimes during a 24 h induction period is reported. There is noevidence that cytokinins act by a ‘triggering’ effectwith a long half life, the response in the target cells beingthe same as that expected from the amount of cytokinin and cytokininmetabolite remaining in the tissue at the time of extraction.Either continuous presence of cytokinin is needed or any triggeraction is short lived, and continuous ‘re-triggering’is needed to achieve maximum response. Key words: Amaranthus tricolor, Betacyanin synthesis, Cytokinin action, Target cells     

Seasonal Changes in the Physiology of S24 Perennial Ryegrass (Lolium perenne L.): 3. Partition of Assimilates between Root and Shoot during the Transition from Vegetative to Reproductive Growth

The partition of ¹⁴C labelled current assimilates to root insimulated swards of Lolium perenne cv. S24 was measured duringthe transition from vegetative growth in autumn to reproductivegrowth in spring under close to natural conditions of lightand temperature. Assimilate partitioning was also measured in‘established’ swards cut three times during thegrowing season and in vegetative ‘seedling’ swardsgrowing in autumn and in spring. All measurements were madewhen the swards had achieved more than 90 per cent light interception,and all swards were abundantly supplied with water and mineralnutrients. During autumn there was a gradual decrease in the proportionof assimilates partitioned to the roots in both the ‘established’and the ‘seedling’ swards. In the established swards,partition to roots was low over winter, increased during earlyspring, but decreased dramatically, later in the spring, whenstem elongation began. In contrast, in the unvernalized vegatativeseedling swards in spring, partition to roots remained high. The seasonal pattern of assimilate partitioning is consideredin relation to changes in the natural environment and the rateat which the crop fixed carbon in photosynthesis. A decreasein the proportion of assimilates partitioned to roots duringlate spring was significant in increasing the production ofshoot at that time but seasonal differences in partition contributedvery little to the marked differences in shoot growth betweenthe spring and autumn crop. Lolium perenne L., perennial ryegrass, partition of assimilates, flowering     

Accumulation of Phenylpropanoids in Cotyledons of Morning Glory (Pharbitis nil) Seedlings during the Induction of Flowering by Poor Nutrition

Flowering of Pharbitis nil strain Violet is induced in continuouslight under poor nutritional conditions. High-performance liquidchromatography of extracts of the cotyledons revealed that twocompounds in addition to chlorogenic acid accumulate under suchconditions. The compounds were identified as pinoresinol glucosideand p-coumaroylquinic acid. The endogenous levels of these phenylpropanoidswere correlated with the flowering response when nutrition waspoor. However, activation of phenylpropanoid biosynthesis seemednot to be essential for the induction of flowering. (Received May 17, 1993; Accepted July 26, 1993)     

Studies on the Role of Photosynthesis in the Photoperiodic Induction of Flowering in the Short-Day Plants Kalanchoe blossfeldiana Poellniz and Xanthium pensylvanicum Wallr: II. THE EFFECT OF CHEMICAL INHIBITORS OF PHOTOSYNTHESIS

The role of photosynthesis in flower induction in the short-dayplants Kalanchoe blossfeldiana and Xanthium pensylvanicum wasinvestigated by chemical suppression of photosynthesis and preventionof chlorophyll formation in the induced leaf. ‘Bleaching’leaves with streptomycin completely prevented flowering in X.pensylvanicum at concentrations shown to reduce the chlorophylland carotenoid content of the leaf significantly. Such leaveswere unable to induce flowering even when supplied with sugarsand other photosynthetic products. Photosystem II inhibitors,DCMU and cadmium ion, inhibited induction in both species aswell as suppressing photosynthesis (as tested by O₂ evolutionand starch production) whereas the photosystem I inhibitor,metronidazole, had no effect. Antimycin A inhibited floweringin K. blossfeldiana and may have a similar site of action toDCMU. Neither ammonium ion nor DBMIB, which acts upon plastoquinone(i.e. between PS I and PS II in the ‘Z scheme’),had any effect on floral induction and it is argued that theinductive process is independent of photosynthetic phosphorylationbut a step in the electron transport pathway between the sitesof action of DCMU and DBMIB may be crucial. DSPD and its hydrolysisproduct, salicylaldehyde, suppressed flowering in K. blossfeldianabut the uncertainty regarding their chemistry precludes anyfirm conclusions regarding the nature of their action.     

Differential Effects of Kinetin on Acidic Ethyl Acetate-soluble and Highly Water-soluble Gibberellin Fractions in 'Single' and 'Double' Nigella damascena L.

The extractable GA-like substances from seedlings of both ‘double’and ‘single’ genotypes of Nigella damascena areequally compartmentalized between AES and HWS fractions. However,when seedlings were floated on an aqueous solution of kinetinprior to extraction, the compart-mentalization profiles werealtered differentially. The levels of the AES fraction from‘double’ tissues increased concomitantly with adecrease in the level of the HWS fraction. No detectable shiftin the activity of AES and HWS fractions was observed in ‘single’seedlings. This differential response to kinetin parallels otherobservations on these two genotypes and substantiates an interpretationthat these two morphologic traits are associated with ratherdifferent capacities to metabolize GAs.     

Changes in Levels of Naturally Occurring Gibberellin-like Substances during Germination of Seed of Lycopersicon esculentum Mill.

Naturally occurring gibberellin-like substances possessing acidic,basic, and neutral properties were detected, by paper partitionchromatography, in ethanolic extracts of tomato seed and ofetiolated seedlings after 72 and 116 hours' growth. Dwarf maizemutants of the d-1 and d-5 types, ‘Meteor’ pea seedlingsand young ‘Potentate’ tomato plants were used asbioassay material. Hydrolysis of seed and seedling proteinsby ficin in phosphate buffer, pH 6.2, after removal of ethanol-solublesubstances, liberated more and different ‘bound’gibberellin-like substances. It is suggested that protein hydrolysisduring germination is an important means of liberating thesesubstances at different stages of seedling development. Acidic substances were present in all the extracts prepared,but in general two with Rfs 0.25 and 0.55 in iso-propanol: ammonia:water : : 10:1:1 v/v were differentiated on d-2 and d-5 maizerespectively. Neutral substances in dry seed extracts chromatographedin the same solvent, had Rfs of 0.05, 0.35, and 0.95 and thesewere found only in the ethanolic (‘free’) extracts.They were active on d-1 and d-2 maize and ‘Meteor’pea. Basic gibberellin-like substances with Rfs of 0.05 and0.35 were found in ‘free’ extracts of both dry seedand etiolated seedlings after 116 hours' growth which were activeon d-2 maize only. Two others with Rfs 0.45 and 0.95 were extractedfrom seedlings after 72 hours' growth and these were activeon young ‘Potentate’ tomato plants. It is suggested that certain gibberellin-like substances, capableof reversing dwarfism in test plants, may be responsible formorphogenetic or other responses not involving stem extensionin the parent species. Changes were found in the levels of gibberellin-likesubstances but there was no evidence of changes in levels ofseed inhibitors relative to seed growth substances.     

Photothermal Time for Flowering in Lentils (Lens culinaris) and the Analysis of Potential Vernalization Responses

Two cultivars of lentils, Laird and Precoz, were subjected to18 potentially vernalizing treatments, comprising constant temperaturesof 1, 5 or 9 °C in factorial combination with photoperiodsof 8 or 16 h for 10, 30 or 60 d. These seeds or seedlings, togetherwith non-vernalized seeds (as controls), were then transferredto four different growing regimes (‘day’/‘night’temperatures of 18/5 °C or 24/13 °C, factorially combinedwith photoperiods of 11 or 16 h). Variation in the number ofdays from sowing to first flower (f) in the growing regimesfor the controls conformed to the equation I/f = a+b+cP, whereis mean temperature (°C), P is photoperiod (h) and a, band c are genotype-specific constants. Accordingly, when theenvironment varies during development, the photothermal timerequired to flower in day-degrees (°C d) is given by 1/babove a base temperature defined as —(a+cP)/b. Most variationin time to flower could be accounted for by the photothermaltime accumulated in the two successive environments. Therefore,there was no evidence of a specific low-temperature vernalizationresponse in either cultivar. Neither was there evidence of ‘short-day’vernalization, i.e. advancement of flowering resulting frompreliminary short-day treatments. A potential error inherentin the predictive model described arises because it ignoresthe presence of a pre-inductive, photoperiod-insensitive phase;but agro-ecological considerations suggest that this error maynot be important in practice. Lens culinaris, lentil, flowering, photoperiodism, vernalization, photothermal time, screening germplasm     

EGTA inhibits floral induction of Pharbitis nil via its influence on gas exchange properties of stomata

The purpose of the study was to determine inhibitory effect of calcium chelator; ethylene glycol-bis(2-aminoethylether)-N,N,N′,N′-tetraacetic acid (EGTA) on flowering of a short-day (SD) plant Pharbitis nil. It was found that 20 mM solution of EGTA applied on cotyledons of 5-d-old P. nil seedlings four hours before the start of 16-h-long induction night decreased the flowering response by 55% compared to the control plants not treated with this Ca²⁺ chelator. It also caused a very significant decrease of photosynthesis rate, transpiration rate and stomatal conductance both in light and darkness conditions. The results of this study confirm earlier hypothesis suggesting the effect of Ca²⁺ and its modulators on P. nil flowering is due to their influence on the stomata.     

Establishment of photoperiodic sensitivity by benzyladenine and a brief red irradiation in dark grown seedlings of Pharbitis nil Chois.

Flowering of etiolated seedlings of Pharbitis nil resulted followinga single, brief red irradiation prior to an inductive dark period.Following this irradiation benzyladenine sprayed on the seedlingsenhanced flowering dramatically and this effect was maximalfor concentrations between 44 and 120µM. In the presenceof benzyladenine a brief (4 to 10 sec) low energy red irradiation(2.6 Wm^–2) resulted in flowering and repeated far-redphotoreversal of this red promotion provided clear evidenceof the sole involvement of phytochrome. However, after suchbrief irradiations the critical dark period for flowering waslonger than is normally found in seedlings grown in light whichindicated that additional photoresponses might be importantin natural conditions. An examination of seedling photosynthesisand assimilate transport indicated that the benzyladenine effecton flowering may relate to its promotion of assimilate and floralstimulus transport to the shoot apex. ¹ Present address: Faculty of Agriculture, Mie University, TsuCity, Mie Prefecture, Japan. (Received August 21, 1978; )     

The Control of Flowering in Wheat and Barley: What Recent Advances in Molecular Genetics Can Reveal

The combined forces of developmental biologists, studying primordiuminitiation at the stem apex, and mathematical modellers, developingsimulations of crop growth and development, have brought aboutconsiderable advances in the understanding of the control offlowering in wheat and barley. Nevertheless, there are stillmajor gaps in this understanding including: what determinesthe basic rate of development (magnitude of the phyllochronor plastochron); how temperature and photoperiod interact tobring about the transition from vegetative to reproductive development;and how flowering occurs eventually in the absence of inductiveconditions. Although geneticists have tended to measure cerealflowering in terms of ‘days from sowing or emergence toheading’, results of studies using aneuploids and molecularmarkers are compatible with the roles for photoperiod and low-temperaturevernalization established in purely-physiological or developmentalinvestigations. They have also revealed the existence of ‘earlinessperse’loci, whose detailed roles have yet to be established.Progress towards isolating and characterizing wheat and barleyloci is hampered by the poor resolution of mapping (locationto a precision of tens of thousands of base pairs). Neitherof these broad approaches promises a rapid resolution of thefactors controlling the induction of flowering. Two expandingareas of molecular genetics now provide potential for greaterunderstanding of cereal flowering. First, the extensive homoeologyamong members of the Gramineae can be employed to establishthe existence and location of genes or quantitative trait lociin rice which correspond to controlling loci in wheat or barley.Since the rice genome is 1/30th of the size of the wheat genome,the accuracy of mapping loci can be much higher, and there isgreater potential for precise location of loci using techniquessuch as chromosome walking. With the ultimate cloning of individualgenes, and the isolation of gene products, the relative rolesof the 20 loci apparently involved in the induction of floweringof wheat could be explored. However, progress in the moleculargenetics ofArabidopsis(the second area) may provide a more rapidroute to understanding the control of flowering in cereals forseveral reasons: its small genome (1/4 that of rice); the likelihoodof extensive homoeology with cereals, in spite of differencesin codon usage between monocots and dicots; the existence ofa wide range of flowering-time mutants; and the control of floralinduction by a similar range of environmental factors includingphotoperiod and low temperature. It is likely that the MCDK(Martinez-Zapater, Coupland, Dean and Koornneef, 1994. In: MeyerowitzEM, Somerville CR.Arabidopsis.New York: Cold Spring Harbor Laboratory,403–433) model, formulated to explain the genetic andenvironmental control of flowering inArabidopsis,could be employedusefully in the formulation of experimental work on floweringin wheat and barley. This paper reviews these issues, payingparticular attention to the significance of ‘earlinessperse’ loci and the ‘constitutive floral pathway’for wheat and barley.Copyright 1998 Annals of Botany Company Wheat, barley, rice,Arabidopsis,flowering, photoperiod, vernalization, genetics, development.     

Developmental Genetics and Morphological Evolution of Flowering Plants, Especially Bladderworts (Utricularia): Fuzzy Arberian Morphology Complements Classical Morphology

This review compares new developmental models on flowering andother vascular plants with evolutionary hypotheses formulatedby Agnes Arber (1879–1960) and like-minded botanists.Special emphasis is laid on philosophical basics such as perspectivism,pluralism about evolutionary modelling, continuum way of thinking,and fuzzy logic. Arber's perspective is best labelled as F uzzyA rberian M orphology (FAM Approach). Its proponents (‘FAMmers’)treat structural categories (e.g. ‘roots’, ‘shoots’,‘stems’, ‘leaves’, ‘stipules’)in vascular plants as concepts with fuzzy borderlines allowingintermediates (including transitional forms, developmental mosaics).The FAM Approach complements Cla ssical Plant M orphology (ClaMApproach), which is the traditional approach in botany. ClaMproponents (‘ClaMmers’) postulate that the structuralcategories of vascular plants are regarded as concepts withclear-cut borderlines and without intermediates. However, duringthe evolution of vascular plants, the root-shoot distinctionand the stem-leaf distinction have become blurred several timesdue to developmental changes, resulting in organs with uniquecombinations of features. This happened, for example, in thebladderworts (Utricularia, Lentibulariaceae). When focusingon the ‘leaf’, the FAM Approach is identical toArber's ‘partial-shoot theory of the leaf’ and Sinha's‘leaf shoot continuum model’. A compound leaf canrepeat the developmental pathway of the whole shoot, at leastto some degree. For example, compound leaves of Chisocheton(Meliaceae)with indeterminate apical growth and three-dimensional branchingmay be seen as developmental mosaics sharing some growth processeswith whole shoots! We focus here on the FAM Approach becausethis perspective is especially promising for developmental geneticistsstudying flowering and other vascular plants. Copyright 2001Annals of Botany Company Review, body plan, developmental mosaics, leaf development, history of botany, homeosis, homeotic genes, Lentibulariaceae, morphological evolution, process morphology, stipules, Utricularia, flowering plants